Continuously missing outer rings in woody plants at their distributional margins

Woody plants growing in seasonal climates normally form one growth ring each year. However, under severe stress conditions they may not complete annual wood production all the way down to the root collar resulting in continuously missing outer rings at lower stem sections (CMORs). Here we test whether CMORs occur at different distributional margins of woody plant species, where stress levels are expected to be high. We tested 13 species (202 individuals) of trees and shrubs growing at elevational and latitudinal shrub- and tree-lines, under conditions that are normally associated with a reduction of radial growth such as drought, cold or nutrient deficiency. Samples were collected from Alaska, north western Russia, Central Europe, Scandinavia and south eastern Spain. Annual radial growth was measured along several disks or cores of each individual applying the serial sectioning technique. Individuals of nine species showed CMORs. The proportion of individuals with CMORs within a sampled species and site ranged from 0% to 80%. The number of CMORs within an individual increased with decreasing sampling height on the stem. Significant correlations existed between the amount of CMORs and (i) cambial age, (ii) stem length, (iii) stem proportion/length below peat surface, and (iv) herbivore-caused defoliation intensity depending on the species and sampling location. Our results suggest that CMORs can be associated with a cambium that may be inactive, yet functional, for up to 18 years. We conclude that CMORs can occur in a wide variety of species, growth forms and biomes, suggesting a general strategy of woody plant growth. To avoid miss-dating in dendrochronological studies, we further recommend the use of the serial sectioning technique for woody plants growing in unfavorable environments, above all, when sample size is limited and when no established chronology for cross-dating is available.